Estimating the body size of eocene primates: A comparison of results from dental and postcranial variables

Estimating body weights for fossil primates is an important step in reconstructing aspects of their behavior and ecology. To date, the body size of Eocene euprimates—the Adapidae and Omomyidae—has been estimated only from molar area. Studies on other primates and mammals demonstrate that body weights estimated from teeth are not always concordant with those estimated from postcranial variables. We derive estimates for Eocene primates based on tarsal bone variables to compare with previously published values derived from dental measures. Stepsirhine-wide, family-level, and subfamily-level models are developed and compared. We also compare the accuracy and precision of dental- and tarsal-based regression models for predicting weight in extant species. Tarsal bone and dental area measures prove to be equally robust in predicting body weight; however, highly disparate estimates are often obtained from different variables. Equations based on lower-level taxonomic groups perform better than more widely based models. However, all equations considered yield fairly large errors, which can affect interpretations of paleoecology. The choice of the more robust prediction is not straightforward.     

Predicting body mass of bonobos (Pan paniscus) with human-based morphometric equations

A primate's body mass covaries with numerous ecological, physiological, and behavioral characteristics. This versatility and potential to provide insight into an animal's life has made body mass prediction a frequent and important objective in paleoanthropology. In hominin paleontology, the most commonly employed body mass prediction equations (BMPEs) are “mechanical” and “morphometric”: uni- or multivariate linear regressions incorporating dimensions of load-bearing skeletal elements and stature and living bi-iliac breadth as predictor variables, respectively. The precision and accuracy of BMPEs are contingent on multiple factors, however, one of the most notable and pervasive potential sources of error is extrapolation beyond the limits of the reference sample. In this study, we use a test sample requiring extrapolation—56 bonobos (Pan paniscus) from the Lola ya Bonobo sanctuary in Kinshasa, Democratic Republic of the Congo—to evaluate the predictive accuracy of human-based morphometric BMPEs. We first assess systemic differences in stature and bi-iliac breadth between humans and bonobos. Due to significant differences in the scaling relationships of body mass and stature between bonobos and humans, we use panel regression to generate a novel BMPE based on living bi-iliac breadth. We then compare the predictive accuracy of two previously published morphometric equations with the novel equation and find that the novel equation predicts bonobo body mass most accurately overall (41 of 56 bonobos predicted within 20% of their observed body mass). The novel BMPE is particularly accurate between 25 and 45 kg. Given differences in limb proportions, pelvic morphology, and body tissue composition between the human reference and bonobo test samples, we find these results promising and evaluate the novel BMPE's potential application to fossil hominins.     

Brief communication: Hair density and body mass in mammals and the evolution of human hairlessness

Humans are unusual among mammals in appearing hairless. Several hypotheses propose explanations for this phenotype, but few data are available to test these hypotheses. To elucidate the evolutionary history of human “hairlessness,” a comparative approach is needed. One previous study on primate hair density concluded that great apes have systematically less dense hair than smaller primates. While there is a negative correlation between body size and hair density, it remains unclear whether great apes have less dense hair than is expected for their body size. To revisit the scaling relationship between hair density and body size in mammals, I compiled data from the literature on 23 primates and 29 nonprimate mammals and conducted Phylogenetic Generalized Least Squares regressions. Among anthropoids, there is a significant negative correlation between hair density and body mass. Chimpanzees display the largest residuals, exhibiting less dense hair than is expected for their body size. There is a negative correlation between hair density and body mass among the broader mammalian sample, although the functional significance of this scaling relationship remains to be tested. Results indicate that all primates, and chimpanzees in particular, are relatively hairless compared to other mammals. This suggests that there may have been selective pressures acting on the ancestor of humans and chimpanzees that led to an initial reduction in hair density. To further understand the evolution of human hairlessness, a systematic study of hair density and physiology in a wide range of species is necessary. Am J Phys Anthropol 152:145–150, 2013. © 2013 Wiley Periodicals, Inc.     

Reproductive strategies,body size,and encephalization in primate evolution

In order to understand fully the generally high level of encephalization observed in living primates, we must determine why early primates exhibited moderately large relative brain sizes compared to their early Tertiary contemporaries. The relatively high degree of encephalization in early primates may be related at least in part to the fact that they were highly unusualamong mammals in combining a small body size with a strongly precocial reporductive strategy. Other small, precocial mammals also exhibit moderately high levels of encephalization; but primates appear to have been truly uniquein being the only such small-sized and highly precocial group to give rise to extensive radiations of larger descendants. This is a key element in understanding primate brain evolution, because the initial “head start” of the early primates was translated up to larger sizes in descendants. The possible relationships among encephalization, precociality, small size, and arboreality are discussed, particularly in light of recent debates concerning the validity of the superorder Archonta. This work emphasizes that we need to consider relative brain size as but one element in a complex synergistic network of morphological and life-history features.     

New body mass estimates for Omomys carteri,a middle Eocene primate from North America

We report new body mass estimates for the North American Eocene primate Omomys carteri. These estimates are based on postcranial measurements and a variety of analytical methods, including bivariate regression, multiple regression, and principal components analysis (PCA). All body mass estimation equations show high coefficients of determination (R²), and some equations exhibit low prediction errors in accuracy tests involving extant species of body size similar to O. carteri. Equations derived from PCA-summarized data and multiple regression generally perform better than those based on single variables. The consensus of estimates and their statistics suggests a body mass range of 170–290 g. This range is similar to previous estimates for this species based on first molar area (Gingerich, J Hum Evol 10:345–374, 1981; Conroy, Int J Primatol 8:115–137, 1987). Am J Phys Anthropol 109:41–52, 1999. © 1999 Wiley-Liss, Inc.     

The scaling of body size and mass in a host-parasitoid association: influence of host species and stage

We describe the allometry of body mass and body size as measured by hind-tibia length in males of Monoctonus paulensis (Ashmead) (Hymenoptera: Braconidae, Aphidiinae), a solitary parasitoid of aphids. To assess the influence of host quality on allometric relationships, we reared parasitoids on second and fourth nymphal instars of four different aphid species, Acyrthosiphon pisum (Harris), Macrosiphum creelii Davis, Myzus persicae (Sulzer) and Sitobion avenae (F.), under controlled conditions in the laboratory. Dry mass was positively correlated with hind-tibia length, and could be predicted from it, in unparasitized aphids, in aphid mummies containing parasitoid pupae, and in the parasitoid. The reduced-major-axis scaling exponents for the regression of dry mass on hind-tibia length were species-specific in aphids, reflecting differences in volume and shape between species. In mummified aphids, the stage at death influenced the size/mass relationship. In males of M. paulensis, the allometric exponent varied between parasitoids developing in different kinds of host. Individuals developing in pea aphid were absolutely larger in dry mass as well as proportionately larger relative to their hind-tibia length. We discuss the allometry of body size and body mass in relation to parasitoid fitness.     

Correlates of sexual dimorphism in primates: Ecological and size variables

The effects of a series of ecological and size factors on the degree of sexual dimorphism in body weight and canine size were studied among subsets of 70 primate species. Variation in body-weight dimorphism can be almost entirely attributed to body weight (83% of variance R² of weight dimorphism). Much smaller amounts of the variation can be attributed to mating system (R² =6.8%,polygynous species being more dimorphic than monogamous ones) and diet (R² = 2.5%,frugivorous species being more dimorphic than folivorous ones). Habitat (arboreal vs. terrestrial) and activity rhythm (nocturnal vs. diurnal) have only an indirect effect on weight dimorphism. Variation in canine-size dimorphism can be explained in terms of canine size (R² =49%),activity rhythm (R² = 20%,diurnal species being more dimorphic than nocturnal ones), and mating system (R² = 10%).Habitat and diet do not play a significant role in canine-size dimorphism. The unexpectedly high contribution of size to sexual dimorphism coupled with the observation of increased sexual dimorphism with increased size leads us to formulate a new selection model for the evolution of sexual dimorphism. We suggest that if there is selection for size increase, whatever its cause, directional selection in both males and females will lead to an increase in sexual dimorphism based on differences in genetic variance between the sexes. Sexual selection, resource division between the sexes, or lopsided reproductive selection need not play a role in such a model.     

Phyletic size change and brain/body allometry: A consideration based on the African pongids and other primates

A problematic aspect of brain/body allometry is the frequency of interspecific series which exhibit allometry coefficients of approximately 0.33. This coefficient is significantly lower than the 0.66 value which is usually taken to be the interspecific norm. A number of explanations have been forwarded to account for this finding. These include (1) intraspecificallometry explanations, (2) nonallometric explanations, and (3) Jerison’s “extraneurons” hypothesis, among others. The African apes, which exhibit a lowered interspecific allometry coefficient, are used here to consider previous explanations. These are found to be inadequate in a number of ways, and an alternative explanation is proposed. This explanation is based on patterns of brain and body size change during ontogeny and phytogeny. It is argued that the interspecific allometry coefficient in African apes parallels the intraspecific one because similar ontogenetic modifications of body growth separate large and small forms along each curve. In both cases, body size differences are produced primarily by growth in later postnatal periods, during which little brain growth occurs. Data on body growth, neonatal scaling, and various lifehistory traits support this explanation. This work extends previous warnings that sizecorrected estimates of relative brain size may not correspond very closely to our understanding of the behavioral capacities of certain species in lineages characterized by rapid change in body size.     

Scaling relationships between craniofacial sexual dimorphism and body mass dimorphism in primates: implications for the fossil record

Craniofacial remains (the most abundant identifiable remains in the fossil record) potentially offer important information about body size dimorphism in extinct species. This study evaluates the scaling relationships between body mass dimorphism and different measures of craniofacial dimorphism, evaluating taxonomic differences in the magnitude and scaling of craniofacial dimorphism across higher taxonomic groups. Data on 40 dimensions from 129 primate species and subspecies demonstrate that few dimensions change proportionally with body mass dimorphism. Primates show general patterns of greater facial vs. neurocranial and orbital dimorphism, and greater dimorphism in lengths as opposed to breadths. Within any species, though, different craniofacial dimensions can yield very different reconstructions of size dimorphism. There are significant taxonomic differences in the relationships between size and craniofacial dimorphism among primate groups that can have a significant impact on reconstructions of body mass dimorphism. Hominoids tend to show lower degrees of facial dimorphism proportional to size dimorphism than other primates. This in turn implies that strong craniofacial dimorphism in Australopithecus africanus could imply very strong body size dimorphism, conflicting with the relatively modest size dimorphism inferred from postcrania. Different methods of estimating the magnitude of size dimorphism from craniofacial measurements yield similar results, and yield comparatively low percent prediction errors for a number of dimensions. However, confidence intervals for most estimates are so large as to render most estimates highly tentative.     

Problems of body-weight estimation in fossil primates

Body-weight estimates of fossil primates are commonly used to infer many important aspects of primate paleobiology, including diet, ecology, and relative encephalization. It is important to examine carefully the methodologies and problems associated with such estimates and the degree to which one can have confidence in them. New regression equations for predicting body weight in fossil primates are given which provide body-weight estimates for most nonhominid primate species in the fossil record. The consequences of using different subgroups (evolutionary “grades”) of primate species to estimate fossil-primate body weights are explored and the implications of these results for interpreting the primate fossil record are discussed. All species (fossil and extant) were separated into the following “grades”: prosimian grade, monkey grade, ape grade, anthropoid grade, and all-primates grade. Regression equations relating lower molar size to body weight for each of these grades were then calculated. In addition, a female-anthropoid grade regression was also calculated for predicting body weight infernales of extinct, sexually dimorphic anthropoid species. These equations were then used to generate the fossil-primate body weights. In many instances, the predicted fossil-primate body weights differ substantially from previous estimates.     

Cranial variables as predictors of hominine body mass

Body mass is a key variable in investigating the evolutionary biology of the hominines (Australopithecus, Paranthropus, and Homo). It is not only closely related to life-history parameters but also provides a necessary baseline for studies of encephalization or megadonty. Body mass estimates are normally based on the postcranial skeleton. However, the majority of hominid fossils are cranio-dental remains that are unassociated with postcranial material. Only rarely can postcranial material be linked with craniodentally defined hominid taxa. This study responds to this problem by evaluating body mass estimates based on 15 cranial variables to determine whether they compare in reliability with estimates determined from postcranial variables. Results establish that some cranial variables, and particularly orbital area, orbital height, and biporionic breadth, are nearly as good mass predictors for hominoids as are some of the best postcranial predictors. For the hominines in particular, orbital height is the cranial variable which produces body mass estimates that are most in line with postcranially generated estimates. Both orbital area and biporionic breadth scale differently in the hominines than they do in the other hominoids. This difference in scaling results in unusually large estimates of body mass based on these variables for the larger-sized hominines, although the three cranial variables produce equivalent predicted masses for the smaller-bodied hominines. © 1994 Wiley-Liss, Inc.     

The relationship between body mass and relative investment in testes mass in cetaceans: Implications for inferring interspecific variations in the extent of sperm competition

Based on sperm competition theory, percentage testes mass (% of total body mass) has been used to infer variations in the extent of sperm competition within mating systems of cetaceans. However, in most amniote taxa, including mammals, there is an underlying negative relationship between body mass and relative investment in testes mass, which must first be taken into account. Here, I identify a very strong nonlinear, negative relationship between body mass in cetaceans and relative investment in testes mass based on data from 31 species. As a result, if percentage testes mass alone is used to infer the relative extent of sperm competition in cetaceans, its importance in mating systems of smaller species is likely to be overestimated, whereas its role in larger species is likely to be underestimated. Similarly, there will also be systematic biases if this relationship is assumed to be linear when it is not. Therefore, it is essential that the underlying, nonlinear body mass–testes mass relationship is correctly taken into account when using relative investment in testes mass to estimate the relative levels of sperm competition in cetaceans. This is particularly important if such inferences are used to inform conservation strategies for endangered cetacean species.     

The relationship between body weight and natural mortality in juvenile and adult fish: a comparison of natural ecosystems and aquaculture

The relationship between body weight and natural mortality in juvenile and adult fish was analysed for different aquatic ecosystems: lakes, rivers, the ocean, and pond, cage and tank aquaculture systems. Mortality was modelled as a power function of weight, and the parameters b (exponent) and M_u (mortality at the unit weight of 1 g) estimated for fish in the six ecosystems, as well as within selected populations, species and families. At the ecosystem level, no significant differences in parameters were found between lakes, rivers and the ocean and a joint mortality-weight relationship for all natural ecosystems was estimated with parameters b=?0.288 (90% CL[?0.315, ?0.261]) and M_u=3.00 (90% CL[2.70, 3.30]) year^?1. Among the culture systems, mortality-weight relationships in ponds and cages were not significantly different and a joint relationship was estimated. The weight exponents of mortality in ponds/cages and tanks were very similar at about b=?0.43, and significantly more negative than in natural ecosystems. Mortalities at unit weight were significantly lower in tanks (0.91 year^?1) than in ponds/cages (2.24 year^?1), and both were significantly lower than in natural ecosystems. No systematic differences were found between the mortality-weight relationships determined for individual populations, species or families, and fish in the respective ecosystems. It is hypothesized that aquaculture mortality-weight relationships indicate the allometric scaling of non-predation mortality, which is therefore more strongly size dependent than predation mortality. If non predation mortality in natural ecosystems shows a similar scaling with body weight, then the allometric exponent of predation mortality must be less negative than that observed for total natural mortality. Implications of the established mortality-weight relationships for aquaculture and culture-based fisheries are discussed.     

Estimating fossil hominin body mass from cranial variables: An assessment using CT data from modern humans of known body mass

Body mass estimates are integral to a wide range of inferences in paleoanthropology. Most techniques employ postcranial elements, but predictive equations based on cranial variables have also been developed. Three studies currently provide regression equations for estimating mass from cranial variables, but none of the equations has been tested on samples of known mass. Nor have the equations been compared to each other in terms of performance. Consequently, this study assessed the performance of existing cranial equations using computed tomography scans from a large, documented sample of modern humans of known body mass. Virtual models of the skull were reconstructed and measured using computer software, and the resulting variables were entered into three sets of published regression equations. Estimated and known body masses were then compared. For most equations, prediction errors were high and few individuals were estimated within ±20% of their known mass. Only one equation satisfied the accuracy criteria. In addition, variables that had been previously argued to be good predictors of mass in hominins, including humans, did not estimate mass reliably. These results have important implications for paleoanthropology. In particular, they emphasize the need to develop new equations for estimating fossil hominin body mass from cranial variables. Am J Phys Anthropol 154:201–214, 2014. © 2014 Wiley Periodicals, Inc.     

Predation by red-jointed fiddler crabs on congeners: interaction between body size and positive allometry of the sexually selected claw

The enlarged (major) claw of male fiddler crabs is used in contestsover breeding burrows and is waved to attract females. We recentlydiscovered that males of the red-jointed fiddler crab, Uca minax,also use the claw to kill smaller-sized fiddler crabs, U. pugnaxand U. pugilator, with which they co-occur in Atlantic coastsalt marshes. Large U. minax males use walking legs or the enlargedclaw to capture prey feeding on moist sand flats. On sand flats,small U. minax males and females are much less common than largemales, suggesting that large males move onto sand flats to seekprey. Males of prey species use the major claw against attackingpredators and, consequently, are more likely than females toescape. In laboratory experiments, large U. minax males weremore likely to attack and kill small-clawed males and femalesthan large-clawed males, consistent with a preference for morevulnerable, less threatening prey. The size of the major clawis a positive allometric function of body size. The allometricfunction varies little among species. Also, the mechanical advantageand indices of closing speed and closing force of the majorclaw, when corrected for body size, are not consistently greaterin U. minax relative to prey species. Thus, predation by U.minax males may reflect the opportunity afforded by larger bodysize and positive allometric growth, which result in a majorclaw that is more massive than the prey it is directed against.     

Locomotion in terrestrial mammals: the influence of body mass, limb length and bone proportions on speed

Traditionally a few limb proportions or total limb lengths have been regarded as indicative of peak running velocity. This is due to physical principles (inferred in- and outvelocities around the joints, stride lengths) and also the observation that fast-moving animals tend to share a number of purported key features which are either absent or not developed to near the same extent in slower moving forms. Previous studies have shown hind limb length and metatarsus/femur ratio to be correlated significantly, albeit modestly with running speed. These studies have nearly all been bivariate analyses. Based on the physical principles, there is reason to suppose that more variables than just m/f ratio could be important as adaptations for fast locomotion, and also that bivariate analyses are too simple. In this study a sample of 76 running mammals was used, with running speeds taken from literature. A number of osteological parameters were discovered to covary significantly with peak running speed, albeit only modestly. Using the information from phylogeny reduced all correlations, often significantly so. Multivariate analyses resulted in markedly higher correlation coefficients. Animals probably do not optimize their anatomy for the purpose of running very fast, which occurs only on rare occasions, but for reducing costs of locomotion.  © 2002 The Linnean Society of London, Zoological Journal of the Linnean Society , 2002, 136 , 685–714.